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Invertebrate Drift studied on Dartmoor: The
"Hidden Harvest"
In 1963 towards the end of an exceptionally cold winter a young man -
JM Elliott - set
off to live on Dartmoor. He chose to camp close to Princetown, the
location of a bleak prison constructed in 1806 to hold French and
American POWs. He was there to collect samples of water from the Walla
Brook - a tributary of the East Dart - as part of his PhD.
What he
discovered was to have a major impact on our understanding of the
interrelationship between insect behaviour and trout feeding patterns.
Elliott anchored a fine mesh net in the Walla Brook, removed it every
three
hours over a 24 hour period and counted the number and types of trapped
insect. He spent 18 months on Dartmoor in order to collect
sufficient
samples to complete this study of insect behaviour.
Elliott discovered that the numbers of insects caught in the net varied
as a function of time of day. He called
this phenomenon "invertebrate drift".
- insects in the 'invertebrate drift' are drifting to
locate fresh
feeding grounds, avoid overcrowding or escape predators
- the term 'invertebrate drift' does not refer to
insects that are ascending to the surface prior to hatching
Elliott's findings generated significant research interest from
freshwater ecologists but are not widely discussed in the British
angling
literature. |
This
graph illustrates 'invertebrate drift' and is based on similar work carried out by the Field
Studies Council (FSC) in late April.
- during the day very few invertebrates were found in
the net
- in the evening - and especially towards
midnight - the numbers increase
- this may be an accidental
consequence of insects
emerging from under rocks to feed and being swept off
- or, it may be a deliberate
attempt by insects to move
to a new area in search of food or to escape from predators
- there is evidence that drift is 'intentional' rather
than 'accidental' behaviour
- the causes of invertebrate drift are still the
subject of academic debate and experimental inverstigation
- the term "behavioural drift" has replaced the
earlier term "invertebrate drift" to emphasise that drifting is part of
the behavioural repetoire of certain insect species
- the term "constant or background drift" refers to
random
accidents, and
- the term "catastrophic drift" refers to insect
movement that occurs when rocks move during
a spate (see Gibbins et al, below)
Interestingly, from an angler's point of view:
- the phenomenon of behavioural drift suggests that
there will be periods during the day when increased insect food is
available to trout in the absence of clearly visible surface activity.
Hence my use of the term "Hidden
Harvest"
- there is a significant correlation between the
tendency of an insect species to engage in behavioural drift and that
species being
eaten by trout (see Rader below)
- smaller insects such as midges (Chironomidae) and
blackflies (Simulidae) drift during the day
- caseless caddis are more likely to drift than cased
caddis
- there is some evidence that younger (small) insects
of a particular species drift during the day, whilst larger individuals
drift at night - this may have implications for hook size
- behavioural drift is higher during the summer than
winter
Ephemeroptera,
Simuliidae, Plecoptera, Diptera and
Trichoptera engage
in
behavioural
drift, for example:
- blue-winged olive nymphs (Baetis)
- caseless caddis (Rhyacophila
and Hydropsyche). Stone cased caddis tend not to drift (weight?)
- chironomids (midges) - note there may be no diurnal
pattern of behavioural drift in many chironomids - it can occur
throughout the day and night (see Moss, below)
- blackfly larvae (Simulium) - but they have a sticky
silk thread
that allows them to return 'home' after drifting
- freshwater shrimp (Gammarus)
These results beg the question "Why don't rivers become empty of
insects as a result of behavioural drift?" Part of the answer may be
that females tend to fly or crawl upstream before depositing their
eggs. Moss
(see below) offers some other explanations and discusses the
possibility that behavioural drift - particularly by larger insects
-may occur more often at night where there is less risk of them being
eaten by
trout.
What
the "L" does this mean for the angler?
The concept of 'behavioural drift' has led me to rethink the way I
imagine underwater insect activity when there is no obvious surface
activity associated with a hatch. I now realise that I was
concentrating too much on a straight
vertical line from river bed to surface.
I was thinking in terms of a
very few dislodged nymphs when there was no visible hatch. Then the
pupa ascending through the water column, breaking through the water
surface, shedding its exoskeleton and emerging as a dun.
Now my thinking is more L
shaped. I have added the horizontal component to represent behavioural
drift.
Behavioural drift explains why it is worthwhile fishing an appropriate
imitation close
to the bed of the river to represent a drifting insect,
especially when there is little surface activity.
This is worth doing during the day because:
- some species - particularly smaller insects and
chironomids - may drift
throughout the day (Allan, 1978)
- there may be fewer
natural insects in the drift to compete with your fly for the trout's
attention
- most importantly, trout expect
to find food drifting close to the bed of the river
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Online resources and references:
- JM Elliott commentary on his paper
"Invertebrate drift in a Dartmoor stream"
- Ed Engle, Fishing Small Flies", Stackpole Books, 2005
- C. N. Gibbins, E. Scott, C. Soulsby & I.
McEwan,The relationship between sediment mobilisation and the entry of
Baetis mayflies
into the water column in a laboratory flume, Hydrobiologia (2005) 533:
115–122. available online
- Moss, Ecology of Fresh Waters:Man and Medium, Past to
Future, page 85, available online
- Ephemeroptera Galactica, Official Web Site of the
Permanent Committee of the International
Conferences on Ephemeroptera, available online
- Rader,
A functional classification of the drift: traits
that influence invertebrate availability to salmonids, Can. J.
Fish. Aquat. Sci. 54(6):
1211–1234 (1997), abstract available online
- Hieber, Robinson and Uehlinger, Seasonal and diel
patterns of invertebrate drift in different alpine stream types.
Freshwater Biology (2003) 48, 1078–1092. available online
- David
Allan, Stream Ecology: Structure and Function of Running
Waters, Kluwer Academic Publishers, 1995
- Siler, Current Research on Macroinvertebrate,
Detritus, and Algal Drift in the Au Sable River, available online
- Svendsen, Quinn, and Kolbe, Review
of Macroinvertebrate Drift in Lotic Ecosystems, available online
- Rydberg, Lecture notes on Stream Drift, available online
- Gary LaFontaine, The Dynamics of Nymph Fishing, Part
3 "What are the trout feeding on?", available online
- Allan,Trout Predation and the
Size Composition of Stream Drift,Limnology
and Oceanography, Vol. 23, No. 6 (Nov., 1978), pp. 1231-1237, available
online
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